DICTYOCRYPHALIDAE Suzuki, 2021

Suzuki, Noritoshi, Caulet, Jean-Pierre & Dumitrica, Paulian, 2021, A new integrated morpho- and molecular systematic classification of Cenozoic radiolarians (Class Polycystinea) - suprageneric taxonomy and logical nomenclatorial acts, Geodiversitas 43 (15), pp. 405-573 : 501-503

Family DICTYOCRYPHALIDAE Suzuki , n. fam.

urn:lsid:zoobank.org:act:C897A495-6E6C-4149-84CE-1324AF0AF58C

Lophophaenida Haeckel, 1882: 430 [nomen dubium, below tribe].

Lithobotryida Haeckel, 1887: 1107, 1111-1112 [nomen dubium, as a family]. — Bütschli 1889: 1983 [as a family]. — Anderson 1983: 29 [as a family].

Lithobotryidae Poche, 1913: 222 [nomen dubium]. — Schröder 1914: 143. — Chediya 1959: 186. — Cachon & Cachon 1985: 295. — Chen et al. 2017: 173.

Lophophaenidae – Campbell 1954: D128 [nomen dubium]. — Petrushevskaya & Kozlova 1972: 534. — Dumitrica 1979: 30. — Blueford 1988: 246. — Nishimura 1990: 85, 87 ( sensu emend. ). — Sugiyama 1993: 51. — Dumitrica 1995: 28. — van de Paverd 1995: 217. — Sugiyama 1998: 233. — De Wever et al. 2001: 224, 226. — Matsuzaki et al. 2015: 42.

Lophophaeninae – Campbell 1954: D128 [nomen dubium]. — Dieci 1964: 187. — Petrushevskaya 1971a: 86-91; 1971b: 989; 1981: 87-88. — Takahashi 1991: 96. — Hollis 1997: 55. — Afanasieva et al. 2005: S292. — Afanasieva & Amon 2006: 139.

TYPE GENUS. — Dictyocryphalus Haeckel, 1887: 1308 [type species by subsequent designation ( Campbell 1954: D128): Cornutella ? obtusa Ehrenberg, 1844a: 77].

INCLUDED GENERA. — Antarctissa Petrushevskaya, 1967: 85 . — Botryopera Haeckel, 1887: 1108 (= Trisulcus synonymized by Petrushevskaya 1975: 591). — Dictyocryphalus Haeckel, 1887: 1308 (=? Cephaluspinus n. syn.). — Nomina dubia. — Lithobotrys, Lophophaena , Lophophaenoma , Lophophaenula .

JUNIOR HOMONYMS. — Dictyocephalus Ehrenberg, 1861 (= Dictyocryphalus, Ehrenberg 1861b ) nec Leidy, 1859; Discocephalus Ehrenberg, 1861 (= Dictyocryphalus, Ehrenberg 1861b ) nec Ehrenberg in Hemprich & Ehrenberg, 1829.

DIAGNOSIS. — Dictyocryphalidae Suzuki , n. fam. are two segment- ed Plagiacanthoidea . The thoracic part with a neck or a suture is generally present. No feet and rarely three spinule-like wing rods from A- and double L-rods are present. The shell is subdivided at variable degrees into a post-lobe on the most apical side of the A-rod, a eucephalic lobe in the space between the A- and V-rods, and an ante-lobe on the more ventral side of the V-rod. In well-developed specimens, the eucephalic lobe is bounded by two arches from both the post- and ante-lobes. The level of the neck and the boundary of the eucephalic lobe are always located above the MB’s level. Both post- and ante-lobes usually develops between the MB and the thoracic part. The cephalic spicular system consists of MB, A-, V-, D-, double L- and Ax-rods. The double l-rod is generally absent. The MB is very short or degrades, becoming a pointed connection (PC) with D- and double L-rods. The PC (or MB) is located in the center of the cephalic cavity. The V-rod is rarely absent at genus or species levels. A basal ring does not exist. Instead, a cephalic basal ring-like structure is connected by the A-rod, V-rod, and several supplemental connecting rods that arise from both the double L-rod and D-rod. Due to the development of basal ring-like structure, true double AL- and double LV-arches are absent. In this case, the arches are coded as A’, double L’, l’-rod. This basal ring-like structure is isolated from the cephalic wall that is joined by many rods around the basal ring. The Ax-rod is very short, except for in the case of Antarctissa . No tubes are found on the cephalis. A-rod is merged to the cephalis. It may also be partly or fully free in the cephalic cavity. If the AL’- and L’V-arches are merged with the shell wall, sutures form on both sides of the eucephalic lobe.

The protoplasm was examined in Dictyocryphalus . The endoplasm is transparent or brown, yields multi nuclei, and occupies the cephalis, and part of the thorax depending on specimens. If present, algal symbionts are scattered outside the shell. No axopodial projection was found so far.

STRATIGRAPHIC OCCURRENCE. — early Middle Eocene-Living.

REMARKS

Except for Dictyocryphalus , the genera of the Dictyocryphalidae Suzuki , n. fam. have stable, cephalic initial spicular and arch system components. The Dictyocryphalidae Suzuki , n. fam. are characterized by the presence of a basal-ring like structure and a retrograding MB or three-pointed AC. Dictyocryphalus has variable systems at the species or infra-species level with the presence of a basal ring that is directly connected with the D- and double L-rods, - the presence of double Al- and double DL-arches, - the presence of double VL-arch instead of DL-arch, - the rare presence of an l-rod, - or the absence of lobes. Some unstable characters found in Dictyocryphalus are also observed in representatives of Dimelissidae . The Dictyocryphalidae Suzuki , n. fam. is easily distinguished from the Pseudodictyophimidae Suzuki , n. fam. by the absence of three feet.

There was some confusion among the genera of the Dictyocryphalidae Suzuki , n. fam., and between the Dimelissidae and the Dictyocryphalidae Suzuki , n. fam. (e.g., Dictyocryphalus vs Lithomelissa ; Botryopera vs Amphimelissa ). Rapid examinations of the presence of a cephalic basal-ring structure and an absence of AL- and DL-arches are unrealistic. The A-rod is not free in the cephalic cavity of Dictyocryphalus whereas it is free in the cephalic cavity of Lithomelissa (Dimelissidae) . Three lobate cephalis are similar in Botryopera and in Amphimelissa (Pylobotrydidae) . Differing from Botryopera , Amphimelissa develops a double l-rod and a multicamerate cephalis that is larger than the thorax. As with the Dimelissidae , all the genus members of the Dictyocryphalidae Suzuki , n. fam. except Antarctissa ( Petrushevskaya 1986) remain unconfirmed by the stratigraphic distribution of these genera and species. Moreover, many genera and species remain undescribed.

The cephalic initial spicular system was documented in Antarctissa ( Petrushevskaya 1986: pl. 1, fig. 9), Botryopera ( Sugiyama 1993: figs 14-17), Dictyocryphalus ( Caulet 1974: pl. 9, figs 4-6?; Nishimura 1990: figs 17.1-17.3, 18.3?; Sugiyama et al. 1992: pl. 16, figs 6, 7; Sugiyama 1993: fig. 23.2; 1994: pl. 5, figs 3, 4?; Funakawa 1994: figs 8.1, 8.2, 8.4; 2000: pl. 1, fig. 4 [wrong plate number is indicated on the true plate 1]; Nishimura & Yamauchi 1984: pl. 32, fi. 6; O’Connor 1997a: pl. 6, figs 6, 7, 8?, 9). Living or protoplasm images were illustrated for Dictyocryphalus ( Matsuoka 1993a: fig. 2.6; Ogane et al. 2010: figs 1.6, 1.7; Suzuki & Aita 2011: fig. 5L; Matsuoka 2017: fig. 22; Matsuoka et al. 2017: appendix B; Zhang et al. 2018: 10, figs 6, 8, p. 19, fig. 28).

The taxonomic validity of the “ Lophophaenidae ” involves very complex problems which include the (A) validity of the type species of Dictyocryphalus Haeckel 1887 ; the (B) validity of the type species of Lophophaena Ehrenberg 1847 ; and (C) the possible designation of a neotype for Lophophaena . The type species of Dictyocryphalus is Cornutella ? obtusa Ehrenberg 1844a designated as such by Campbell (1954: D128) as an objective synonym of Dictyocephalus . The name-bearing specimen was first published by Ehrenberg (1854c: pl. 22, fig. 40). The type locality of D. obtusus (Ehrenberg) is Caltanisetta, West of Sicily ( Ehrenberg 1844a: 77), and thus the type specimens were expected to be preserved in Ehrenberg’s slide tray K28B06 ( Suzuki et al. 2009c: 88) in the Ehrenberg collection. The slide series of “Caltanisetta” (K28B06) was highly damaged and many slides of the Caltanisetta are missing. For this reason, Suzuki et al. (2009c) examined all the pieces of the slides, including two-millimeter fragments, and took photographs of all the encountered radiolarian specimens, published in pls 1-21 of Suzuki et al. (2009c). Following this observation, the type specimens appear to be completely missing. Instead, the most similar morphotypes are illustrated in pl. 20, figs 13b-14 of Suzuki et al. (2009c), but their designation of neotype was unlikely because the slides in K28B06 almost completely missing. No raw samples are archived in NfM. The sample locality information is noted in Ehrenberg (1839: 78), but the specification of the locality was unhelpful. Some papers illustrated radiolarians from Sicily ( Riedel & Sanfilippo 1978b; Sanfilippo et al. 1978, 1985; Cortese & BjØrklund 1999). The morphotype that most closely resembles D. obtusus from Caltanisetta was illustrated in Cortese & BjØrklund (1999: figs 21.P-21.R). If we compare pls 1-21 of Suzuki et al. (2009c) with the figures 20-22 of Cortese & BjØrklund (1999), the fauna appears to be nearly identical, and it may be tentatively concluded that figs 21.P-21.R of Cortese & BjØrklund (1999) represent the true D. obutsus as a potential neotype.

A second encountered problem is the type species of Lophophaena . The genus Lophophaena was established by Ehrenberg (1847) without any included species, and the first assigned species was “ Lophophaena Galea Orci ” as a monotype ( Ehrenberg 1854b: 245). Thus, this species automatically becomes the type species of Lophophaena . The name-bearing specimen was noted as “Ex abysso 12000 ped” in the description ( Ehrenberg 1854b), meaning “from 12,000 fathoms in deep”. The fact that the sample information written in Ehrenberg (1854a: table) is noted as 8160’ and not 12000’ is bizarre. The mismatch of type locality is another new problem. Putting aside a sample mismatch, the exact sample locality for “ Lophophaena Galea Orci ” are the samples from “ 42 41’N, 24 35’W, 18 July, 1360 Fath-6480’” or “ 54 17’N, 22 33’W 22 Aug. 2000 Fath-12000’” ( Ehrenberg 1854a: 60). Based on these disparate localities, a new problem arises.Following these papers as well as the internal documents in NfM, potential type series could be found in “Meersgrund II (K27B02)” or “Meersgrund II (K27B03)” ( Suzuki et al. 2009c: 90). All the specimens assigned by Ehrenberg himself are photographed on pl. 30 for K27B02 and pls 31-36 for K27B03 in Suzuki et al. (2009c). The exact sample information is specified on these trays (K27B02 and K27B03). Congruently, any typebearing specimens for “ Lophophaena Galea Orci ” could not be found in the slides. An additional problem stems from the publication. Campbell (1954: D128) falsely indicates “ Lophophaena galea Ehrenberg, 1854a ” as the subsequent type species of Lophophaena . This species has not been formally described and is therefore a nomen nudum. Some papers cite pl. 8, figs 12 of Ehrenberg (1876) as “ Lophophaena galea ” but the name on the plate explanation is “ Lophophaena ? galeata ” which was first described by Ehrenberg (1874: 242-243). In addition, the specimen illustrated in Ehrenberg (1876: pl. 8, fig. 12) is from Barbados and it is not from the true locality of “ L. Galea Orci.” Thus, the correct name-bearing specimen has not been illustrated and has not been preserved in the Ehrenberg collection, resulting in the assigned status as nomen dubium.

VALIDITY OF GENERA

Dictyocryphalus

The translated diagnosis from the original Spanish for Cephaluspinus follows. “ Shell campanulate sub-divided into cephalis and thorax. Surface perforated by sub-circular pores of different size and irregularly distributed. Cephalis with many spines, some larger than others, some of which are apparently broken in the analyzed specimen. The other part of the shell is smooth, except for the basal part, which has spines, or feet, that are in fact externally prolonged terminations of the pore frames. These feet are numerous, approximately 20, and shorter than the cephalic spines. ” Petrushevskaya (1981: 90) considered Cephaluspinus a subjective synonym of Lophophaena because its morphological characteristics correspond entirely to that of Lophophaena . As Lophophaena (sensu Petrushevskaya 1981) corresponds to Dictyocryphalus under the strict ruling under the Code (see the remarks in the Dictyocryphalidae Suzuki , n. fam.), the synonymy has been simply replaced by Dictyocryphalus . However, this synonymy cannot be precisely determined because the initial spicular system of Cephaluspinus is unknown.